After the second Henn et al. paper I did download the data. Unfortunately there are only 62,000 SNPs intersecting with the HGDP. This is somewhat marginal for fine-grained ADMIXTURE analyses, though sufficient for PCA from what I recall. That being said, the intersection with the HapMap data sets runs from ~190,000 SNPs, to the full 250,000 SNPs (this makes sense since the Henn et al. #2 data set has some HapMap populations in it). So I’ve been experimenting a fair amount in the past few days, and I thought I would post on one issue which was clear in the original paper, but which I have replicated.

The Fulani (Fula) people of the western Sahel seem to have a relatively old West Eurasian component which has distinct affinities with the “Maghrebi” element discerned by Henn et al. In fact, the non-Sub-Saharan African ancestry of the Fulani is almost exclusively of this origin. To me this serves as a peculiar mirror of what you see in the Cushitic and Ethiopian Semitic peoples of the far east of the Sahel-Sudan latitudinal region. These populations also seem to be compounds of a Sub-Saharan Africa element with a West Eurasian one, but in their case the admixture is almost exclusively from a Southwest Eurasian (Arabian) component. Geographically these two symmetric admixture events make sense, but the exclusivity is still a bit surprising. Additionally, in both the case of the Fulani and the Ethiopian and Cushitic groups the admixture is widely distributed and even enough to imply that they are old events. I also assumed this because in some admixture runs a “pure” Fulani cluster partitions out, which is not unexpected for stabilized hybrid populations (all human populations are stabilized hybrids if you go back far enough).

To give you a flavor of what I’m talking about here are some screen shots of a run which is currently going. It has 180,000 markers. I removed Tunisians and many African populations from the Henn et al. data set, and included in the Utah whites from the HapMap. The individual plots show the ancestral proportions for each Fulani in the data set:

So what can we see here? First, let’s reiterate something: as in the case of the populations of the Horn of Africa the West Eurasian element in the Fulani is difficult to find in “pure” form in the populations from which it putatively derived. What does that imply? I think that that means that the Fulani have an origin in relatively recent historic time, on the order of 2,000, not 10,000, years. That is because I am skeptical that the Fulani would be able to maintain genetic distinctiveness for ~10,000 years from other populations around them. In contrast, the last 2,000 years have seen the rise of various cultural institutions, from trans-Saharan nomadism to Islam, which might slow down admixture sufficiently to maintain the differences between the Fulani and their neighbors. It also implies to me that the non-Maghrebi “Near Eastern” element which Henn et al. discerned is relatively a recent phenomenon in northwest Africa, else the Fulani should also carry it. How recent? Probably from Classical Antiquity down to the Muslim period. Observe that many North Africa groups have a red “European” element. This may be from Near Eastern populations, but I suspect that the fraction here is just too high to be explained by that. Also, you can see above that some groups in Morocco have nearly as much of this as Egyptians, but far less of the more genuine Near Eastern components.

In all likelihood the West Eurasian component came to the Fulani via the Tuareg or a related or antecedent population. So if you typed the Tuareg you would probably get a better sense of the “pure” “Maghrebi” genetic profile. These genetic results also can serve as fodder to understanding the ethnogenesis of the landscape of the Sahel. In the map above it is interesting to observe that the Hausa speak an Afro-Asiatic language, even though their West Eurasian component is far lower than the Fulani, who speak Niger-Congo dialects. What gives? I suspect that the difference here is that the Hausa are a case of elite emulation of a cultural complex which was much more integrated and elaborated by the time it arrived on the West African scene. This explains how there could be language shift, while in the case of the Fulani there was none. Another hypothesis is that Afro-Asiatic derives from Sub-Saharan Africa itself, and the Chadic (Hausa) group are basal to the phylogeny. I’ll let readers explore the implications of that. A final aspect, I put the quotations in the title because perhaps the Berber dialects spread via elite emulation, and the original Maghrebi ancestors of the Fulani spoke a different language, which has been lost? As they say, for every answer there bloom a thousand questions….

Just to clarify, the Fulanis in question are Wodaabe nomads from the vicinity of Lake Chad, these nomads were sampled in Nigeria but are geographically wide spread (they’re found in Nigeria, Niger, Chad, northwestern Central African Republic, and northern Cameroon… of shoots include the Sudanese Fulani) in addition to being numerically small and demographically insignificant. These Fulanis are not equivalent to the Fulani of the Hausa-Fulani ethnicity in Nigeria (whom happen to be more numerically Hausa with a rather small Fulani sub-group); The probability of significant NW African ancestry amongst these Fulanis is significantly decreased in comparison to the Wodaabe who likely had more contact with groups like the Tuareg and have remained relatively isolated after the admixture event. Anyways, I’m assuming that such a NW African affinity among the Wodaabe (and to an extension Eastern “Chadic” Fulanis) isn’t exactly comparable to the biological affinities of the Western Fulani (the majority) in places like Senegal and Guinea. The Wodaabe, being a relatively small group, likely absorbed admixture from the Tuareg during their relatively historic expansion across the Sahelian region; this is probable, in contrast to some sort of ancient admixture event in the vicinity of Senegal which somehow had no genetic implications for neighboring ethnic groups like the Mandinka (who live in close quarters to the Fulani) and other Senegambian populations. The Wodaabe are only about ~50,000 people (the larger Western Fulani sub-group, i.e. Fulanis from Nigeria, Niger, Chad, the Central African Republic, and Cameroon, are only about ~5,000,000 people) so an admixture event with the larger Tuareg population in the region is the much more likely scenario.

I also wouldn’t be to fast to generalize the “Eurasian” affinity among either the Wodaabe or NE Africans as direct gene-flow via Eurasia. While theirs definitely some real Eurasian admixture in the northern Horn, i.e. Eritrea and the Ethiopian provinces of Tigray and Amhara in particular (in addition to N.Sudan due to more historic events), the Eurasian affinity in the Horn of Africa and in extension SE Africa doesn’t necessarily equate to Eurasian gene-flow and likely imo encompasses indigenous NE African diversity; the Somali and other Eastern Cushities are likely legitimate proxies to any ancestral NE African population given that they lack any additional Eurasian and divergent African admixture as is the case in places like the northern Horn and/or SE Africa respectively. In the case of this pristine NW African cluster in the Fulani, a partial NE African would explain some of it’s affinity to Eurasia. Further study is necessary for a concrete conclusion or rather in establishing a consensus.

http://blogs.discovermagazine.com/gnxp Razib Khan

The Wodaabe, being a relatively small group, likely absorbed admixture from the Tuareg during their relatively historic expansion across the Sahelian region;

when was this? the lack of variance is indicates something older than the past few hundred years for sure.

the Somali and other Eastern Cushities are likely legitimate proxies to any ancestral NE African population given that they lack any additional Eurasian and divergent African admixture as is the case in places like the northern Horn and/or SE Africa respectively.

you need to outline your model. the Fst between the sw asian and other “eurasian” clusters is not very high. i want specific dates you have in mind. e.g., i can’t believe that the “sw asian” in somalis (who seem to lack a “semitic component” as per my earlier post, which is find on tigray, etc.) separated 50,000 years BP from other eurasians. the Fst is too small, and it looks like it is in the west eurasian clade, not basal to other eurasian clades.

Lank

One note, Razib. I think you should have included a proper West African reference for the Fulani. Other ADMIXTURE runs have shown that the West African ancestry of the Fulani is almost exclusively from a ‘northern’ West African source, similar to the Mandenka or Dogon. This makes sense as the Fula language is closely related to Senegambian Niger-Congo languages.

The Bulala, a Nilo-Saharan people, are a poor proxy for this West African ancestral population. Mandenka, Yoruba, or even Bantu would work better. Actually, the slightly increased Eurasian affinity of the Bulala relative to West Africans (present in all East African-derived populations) probably results in a slightly lower North African contribution to the Fulani gene pool.

I don’t believe that the lack of Near Eastern in the Fulani is indicative of recent admixture in Northwest Africa. Many North Africans are similar to East Africans in the sense that their Near Eastern element is often exclusively associated with Southwest Asians. This is visible in Dodecad, among other analyses. There is often a remarkable lack of “Caucasus” or “West Asian” admixture in the Maghreb, which I believe would not be the case if they had non-negligible Arab admixture.

Rather, I hypothesize that the lack of Near Eastern in the Fulani is because their North African ancestors are from areas deep into the Sahara desert, where the people carried a very distinctly “North African” genetic signature (the component that peaks in Mozabites or Sahrawis so far).

Eze

”One note, Razib. I think you should have included a proper West African reference for the Fulani. Other ADMIXTURE runs have shown that the West African ancestry of the Fulani is almost exclusively from a ‘northern’ West African source, similar to the Mandenka or Dogon. This makes sense as the Fula language is closely related to Senegambian Niger-Congo languages.”

Fulanis seem to be mostly of Senegambian origin with Berber/Saharan admixture.

Lank

@Eze,

I agree.

idurar

I suggest you to remove southern Moroccans (heterogenous group with clearly a majority of recently SSA-admixed individuals) and to remove Sahrawis who, like Mozabites, seem to skew the results:
it makes no sense for Egyptians to be more ‘european’ than Berbers (lol)

The Sahrawi-centered component is useless.

http://washparkprophet.blogspot.com ohwilleke

In the uniparental studies I’ve seen, Hasua Y-DNA (which is heavy in R1b-V88 that is present only at very low frequencies or absent in many neighboring populations regardless of linguistic affiliation) is much more uniform than that of the Fulani which shows a greater mix of haplogroups. This would be consistent with a male dominanted elite of more recent origins of than the Fulani.

I would be cautious in describing Chadic languages as basal in Afro-Asiatic. It is basal only in the sense that we are sure it is Afro-Asiatic and aren’t sure how it is related to other Afro-Asiatic language families, not in the sense that it is a likely source language of Afro-Asiatic language speakers from which other Afro-Asiatic language families derived.

The position of the major language families of Afro-Asiatic (Chadic, Cushitic, Berber, Coptic, Omotic and Semitic) is hotly disputed, with almost every arrangement having some legitimate linguist championing it. It is safe to say that Ethiosemetic languages followed Cushitic languages in Ethiopia from a Levantine source, but is hard to say much more, and there is no one genetic signal that really consistently unifies all of the peoples who speak Afro-Asiatic languages in uniparental or autosomal genetics. The lack of anything even as distinct as the sometimes hard to pin down genetic markers of the Turkic or Indo-European languages suggests that some of the major language families in the Afro-Asiatic grouping acquired the language almost entirely via language shift rather than population replacement or admixture (a la Hungarian in Hungary).

For example, R1b-V88, which is a classic signal of Chadic populations, is almost absent from several other Afro-Asiatic language speaking populations (e.g. Berbers) who are geographically close, and has its closest phylogenetic neighbors in remote places like Central Asia and the Dead Sea, while characteristic lineages and genetic markers of Berbers, in turn, are uncommon in in the Hasua.

If I had to guess which of the Afro-Asiatic major language families I was most sure was not an original source for other Afro-Asiatic languages, I’d probably pick the Chadic languages, because the people who speak them are more cleanly distinct genetically than some of the alternatives. Cushitic, Coptic or Semitic branches of the Afro-Asiatic language family seem likely to be more closely related to the proto-language of the macro-language family.

My own personal intuition is that Afro-Asiatic languages have roots in the diffusion of the Neolithic, at least from pre-dynastic Egypt and possibly all the way back to early Jericho, with Chadic and Ethiosemitic languages representing later waves of migration and Omotic reprsenting strong substrate influence or creolization from outside the Afro-Asiatic language family, while I’d associate pre-Bantu Niger-Congo languages with the expansion of Sahel agriculture with a different package of crops, whenever that happened. But, it isn’t something I’d bet money on.

The Mandenka, Wolof, Dogon and Fula languages don’t seem to have as many of the really distinctive traits of “core” Niger-Congo languages in phoetics or grammer, in a pattern rather similar to that of Swahili relative to other Niger-Congo languages, which we know to have historically been a lingua franca acquired by many speakers as adults and under some creole pressure from Arabic. This seems consistent with these languages being under linguistic pressure and influence from Berber languages (and perhaps later Arabic) from the North, with admixture being likely as well.

Eurologist

“My own personal intuition is that Afro-Asiatic languages have roots in the diffusion of the Neolithic”

I think that was likely the latest part of several waves. During LGM, the Gulf of Suez largely did not exist, and the Red Sea was much smaller. Given the great diversity of Afro-Asiatic in that region, I think it is reasonable to assume that proto-AA originated then and there. Before the first waves of agriculturalist, there was a wave of (largely fishing and hunting based) expansion during the Green Sahara, starting about 9,000 years ago, which followed the reappearance of the Red Sea as an obstacle. I envision a two-fold split and expansion during this time: a direct western expansion through Sudan and west, and the emergence of Egyptian/Semitic/Berber in the Northern Nile/Suez region, followed by a westward expansion along the Mediterranean and inland (Berber). This last one might already have involved part of the more typical neolithic package, or more likely, there were two waves in short succession.

By 6,000 ya, climate became much dryer (similar to today), and the split into Semitic might date to that, when travel across the desert became increasingly difficult. This is also consistent with modern computed tree and Bayesian analyses of Semitic.

http://blogs.discovermagazine.com/gnxp Razib Khan

re: fulani reference. i’m not super interested in ss-african ancestry in these runs, so that’s why i didn’t include them. i do have them in other runs, and the same problem crops up where ss-africans “hog up” all the ADMIXTURE components

Onur

it makes no sense for Egyptians to be more ‘european’ than Berbers (lol)

It makes perfect sense given the geography. The Mediterranean Sea has been a very strong barrier to gene flow.

Eze

”it makes no sense for Egyptians to be more ‘european’ than Berbers”

There were no northern West Asian (Caucasus region) groups used here. If you look at the Eurasia7 run of Dodecad you can see that Egyptians indeed have less European but more West Asian admixture than Berbers.

Egyptians have 6.4% Atlantic_Baltic and 28% West Asian, while Mozabites have 21.2% Atlantic_Baltic and only a mere 0.4% West Asian. This also proves that Mozabites weren’t affected by the Arab expansion.

http://blogs.discovermagazine.com/gnxp Razib Khan

yes, west asian would be “european” in this run to a large extent. the Fst between west asian and northern european are usually the smallest pairwise for west eurasian groups.

http://lughat.blogspot.com L

A Berber influence on Fulani has long been suggested by historians; it’s nice to see the genetic data corroborating it. However, this influence is unlikely to be Tuareg. Fulani originated around northern Senegal; until its relatively late eastward expansion, the only branch of Berber in a position to influence it would be Zenaga, which happens to be the single most divergent Berber language. All but a few thousand Zenaga speakers have long since shifted to Arabic, but the Sahrawi sample likely includes some of their descendants, and a Mauritanian sample would make an even better point of comparison.

“perhaps… the original Maghrebi ancestors of the Fulani spoke a different language, which has been lost?” – Unlikely. As noted by Faidherbe (1882:31), the Fulani word for “hundred”, teemedere, is a loanword from Berber, as is the word for “camel” (more recently discussed by Kossmann in Berber Loanwords in Hausa); so we know the early Fulani were in contact with Berber speakers.

Abdisamad

Salam, Can anyone tell me an approximate population of the Fula people in Africa or World wide?

http://washparkprophet.blogspot.com ohwilleke

@14 On the order of 13 million,basically in and around the African Sahel, broken down, per Wikipedia by divisions within the language subfamily:

Of course, definitional issues and measurement accuracies put large error bars around all of these numbers that I wouldn’t trust to be consistently more spot on than a factor of two from the true value. So far as I know, there is not a very large Fula diaspora, although if there is any, it would probably be in urban areas of France which was the former colonial power in much of the Fula language speaking world.

A particularly important definitional issue is an increasing sense of ethnic solidarity between Hasua and Fulani peoples.

The Fulani are one of the numerically larger ethnicities in Africa, but like the Kurds, are cursed with populations that aren’t a good fit to colonially established national boundaries.

Grey

“it makes no sense for Egyptians to be more ‘european’ than Berbers”

It depends what you’re measuring. If you had a population from where segments moved to both Egypt and Europe then you might be measuring what was originally pre-european. This might be partially disguised if the original region and the terriotory between it and Egypt had been partly overrun in the intervening millenia by other peoples.

For example, imagine orange paint from Anatolia to Greece and Anatolia through the Levant to Egypt being then half-covered over with purple paint from Arabia to Egypt and Arabia to the Levant.

http://forwhattheywereweare.blogspot.com/ Maju

When, back in December, I performed almost the same (but not quite identical) analysis as Henn and co. had probably already finished in their lab (but was unpublished, so I did not know) I found that the Fulani do appear to have a small but notable West Eurasian affinity that resolved first into a “Sahrawi” component since K=4. However, at K=8 and then again since K=10, the West Eurasian affinity vanished for a Fulani-specific component instead (plus small Mandenka affinity).

I was rather pissed off because I was trying to figure out North Africans, not Fulbe and there was no apparent Fulbe ancestry in North Africans (Mandenka there was some). But still the result has some interest.

But the really interesting thing is that the Fulani-specific component is equidistant at a relatively low Fst from the Mandenka and Sahrawi components (the latter clearly akin to West Eurasians but the former not at all), what seems to confirm that there is some very old admixture that has been “homogenized” within the Fulani ethnicity.

I would indeed have preferred genuine Fula from Futa Toro or Futa Djallon, who are the original Fulani or Peul population by all accounts but I have no reason to think this should not apply to all or most Fulani in general.

A question remaining is how old is “very old”…

Doctoris Scientia

Originally posted by Razib Khan:
“when was this? the lack of variance is indicates something older than the past few hundred years for sure.”

I’m currently using a modem system to connect to the internet so I’m only going to be able to answer the first half of your question at this particular point in time (since it’s on topic), my apologies, but once I’m back in Cairo I’ll make sure to quickly respond to the entirety of your post.

Not necessarily! An ancient time depth in reference to the admixture date/event doesn’t necessarily explain the aforementioned trend regarding the “NW African” affinity among the Wodaabe Fulani, or at least it’s not the only possibility. An ethnic unit, with a relatively small population size, absorbing indiscriminate gene-flow from another group would likely result in a relatively even distribution of the aforesaid admixture, no matter the time depth; for example, SE African Nilo-Saharan speakers, including the likes of the Maasai and Samburu, are relatively recent migrants, past couple hundred years, from the direction of S. Sudan yet possess a rather evenly distributed of NE African and Bantu admixture. The distribution of the “NW African” component among the Wodaabe simply rules out the possibility of discriminate factors, i.e. slavery, in the facilitation of “NW African” gene-flow into this particular population.

Btw, they do show some variation among themselves and according to Tishkoff et al (2007) the settled Nigerian Fulani possessed less of the “Fulani” cluster than the Wodaabe, ~40 vs. ~60-70%.

Originally posted by L:
“However, this influence is unlikely to be Tuareg. Fulani originated around northern Senegal; until its relatively late eastward expansion, the only branch of Berber in a position to influence it would be Zenaga, which happens to be the single most divergent Berber language.”

While the only way of confirming the likely origins of the “NW African” affinities among this particular West African group is to test other Fulani groups from the vicinity of Senegal and so fourth, the Tuareg origin of the “NW African” affinity among the Wodaabe Fulani remains a plausible explanation. If the Fulani as an entire group, and not particularly the Wodaaabe and to an extent the Eastern Fulani in general (the Fulani individuals sampled by Henn were all Wodaabe), acquired a “NW African” affinity from an ancient event with the Zenega or any other similar group we’re going to have to explain the lack of such admixture among closely related (biocultural, linguistic, geographic etc) groups like the Mandinka for example. On the other hand the Eastern Fulani were rather an exception in their expansion across the Sahel/Sahara and the absorption of various local groups, including possibly the Tuareg; this scenario would explain the uniqueness of the Wodaabe and other Fulani groups in their possession of “NW African” admixture in West Africa, outside Berber speaking groups like the Tuareg.

It’s not uncommon for West African languages, especially those spoken by predominantly Muslim ethnic groups, to have loan words from either Berber or Arabic sources; in particular terms in reference to such things as the camel or numeric values. The only West African language with a genuine Berber sub-structure is northern Songhai in southern Algeria, which isn’t exactly West African in that case.

@ ohwilleke

What also needs to be taken into consideration on the basis of further academic study regarding the Fulani is that the Eastern Fulani often include many outliers due to their expansionist past into the region; for example, half of the Cameroonian Fulani from Tishkoff et al (2009) lacked the “Fulani” cluster and therefore the “NW African” affinity all together and were practically identical to nearby Chadic speakers, while the remaining half were no different from the Wodaabe. The Western Fulani are likely more homogeneous due their stable and constant presence in the greater Senegambian region.

pconroy

@7,

Yeah R1b-V88 is fascinating, as I know a number of Ashkenazi Jews who carry it, so it could be said to be linked to Semitic speakers. I believe Jerico has > 30% R1b-V88. Other branches of R1b exist also in Jewish groups.

It would seem to me that at one time there was quite a lot of R1b in the area of Southern Armenia, Northern Syria and Lebanon and South Central Turkey – an area today where groups like the Alawites, Assyrians still exist and carry higher R1b levels than their neighbors.

As I’ve mentioned previously, it would seem that this area was the staging ground for cattle herders to spread from in many directions. I think Y-DNA J1 was a later entrant to the area from the Caucasus, and eliminated much of the original R1b.

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About Razib Khan

I have degrees in biology and biochemistry, a passion for genetics, history, and philosophy, and shrimp is my favorite food. In relation to nationality I'm a American Northwesterner, in politics I'm a reactionary, and as for religion I have none (I'm an atheist). If you want to know more, see the links at http://www.razib.com